High Stressful Temperature and Genetic Variation of Five Quantitative Traits in Drosophila Melanogaster

Variation of five quantitative traits (thorax length, wing length, sternopleural bristle number, developmental time and larva-to-adult viability) was studied in Drosophila melanogaster reared at standard (25°C) and high stressful (32°C) temperatures using half-sib analysis. In all traits, both phenotypic and environmental variances increased at 32°C. For genetic variances, only two statistically significant differences between temperature treatments were found: the among-sire variance of viability and the among-dam variance of developmental time were higher under stress. Among-sire genetic variances and evolvabilities were generally higher at 32°C but narrow sense heritabilities were not. The results of the present work considered in the context of other studies in D. melanogaster indicate different patterns of genetic variation between stressful and nonstressful environments for the traits examined. Data on thorax length and viability agree with the hypothesis that genetic variance can be increased under extreme environmental conditions. This revised version was published online in July 2006 with corrections to the Cover Date.     

Genetic and phenotypic relationships between immune defense,melanism and life-history traits at different temperatures and sexes in Tenebrio molitor

Insect cuticle melanism is linked to a number of life-history traits, and a positive relationship is hypothesized between melanism and the strength of immune defense. In this study, the phenotypic and genetic relationships between cuticular melanization, innate immune defense, individual development time and body size were studied in the mealworm beetle (Tenebrio molitor) using three different temperatures with a half-sib breeding design. Both innate immune defense and cuticle darkness were higher in females than males, and a positive correlation between the traits was found at the lowest temperature. The effect of temperature on all the measured traits was strong, with encapsulation ability and development time decreasing and cuticle darkness increasing with a rise in temperature, and body size showing a curved response. The analysis showed a highly integrated system sensitive to environmental change involving physiological, morphological and life-history traits.     

Effect of low stressful temperature on genetic variation of five quantitative traits in Drosophila melanogaster

A half-sib analysis was used to investigate genetic variation for three morphological traits (thorax length, wing length and sternopleural bristle number) and two life-history traits (developmental time and larva-to-adult viability) in Drosophila melanogaster reared at a standard (25 degrees C) and a low stressful (13 degrees C) temperature. Both phenotypic and environmental variation showed a significant increase under stressful conditions in all traits. For estimates of genetic variation, no statistically significant differences were found between the two environments. Narrow heritabilities tended to be higher at 13 degrees C for sternopleural bristle number and viability and at 25 degrees C for wing length and developmental time, whereas thorax length did not show any trend. However, the pattern of genetic variances and evolvability indices (coefficient of genetic variation and evolvability), considered in the context of literature evidence, indicated the possibility of an increase in additive genetic variation for the morphological traits and viability and in nonadditive genetic variation for developmental time. The data suggest that the effect of stressful temperature may be trait-specific and this warns against generalizations about the behaviour of genetic variation under extreme conditions.     

Altitudinal patterns for longevity, fecundity and senescence in Drosophila buzzatii

We tested for variation in longevity, senescence rate and early fecundity of Drosophila buzzatii along an elevational transect in Argentina, using laboratory-reared flies in laboratory tests performed to avoid extrinsic mortality. At 25 °C, females from lowland populations lived longer and had a lower demographic rate of senescence than females from highland populations. Minimal instead of maximal temperature at the sites of origin of population best predicted this cline. A very different pattern was found at higher test temperature. At 29.5 °C, longevity of males increased with altitude of origin of population. No clinal trend was apparent for longevity of females at 29.5 °C. There was evidence for a trade-off between early fecundity and longevity at non-stressful temperature (25 °C) along the altitudinal gradient. This trait association is consistent with evolutionary theories of aging. Population-by-temperature and sex-by-temperature interactions indicate that senescence patterns are expressed in environment specific ways.     

Temperature fluctuations during development reduce male fitness and may limit adaptive potential in tropical rainforest Drosophila

Understanding the potential for organisms to tolerate thermal stress through physiological or evolutionary responses is crucial given rapid climate change. Although climate models predict increases in both temperature mean and variance, such tolerances are typically assessed under constant conditions. We tested the effects of temperature variability during development on male fitness in the rainforest fly Drosophila birchii, by simulating thermal variation typical of the warm and cool margins of its elevational distribution, and estimated heritabilities and genetic correlations of fitness traits. Reproductive success was reduced for males reared in warm (mean 24 °C) fluctuating (±3 °C) vs. constant conditions but not in cool fluctuating conditions (mean 17 °C), although fluctuations reduced body size at both temperatures. Male reproductive success under warm fluctuating conditions was similar to that at constant 27 °C, indicating that briefly exceeding critical thermal limits has similar fitness costs to continuously stressful conditions. There was substantial heritable variation in all traits. However, reproductive success traits showed no genetic correlation between treatments reflecting temperature variation at elevational extremes, which may constrain evolutionary responses at these ecological margins. Our data suggest that even small increases in temperature variability will threaten tropical ectotherms living close to their upper thermal limits, both through direct effects on fitness and by limiting their adaptive potential.     

The influence of maternal thermal environments on reproductive traits and hatchling traits in a Lacertid lizard,Takydromus septentrionalis

The thermal environment can induce substantial variation in important life-history traits. Experimental manipulation of the thermal environment can help researchers determine the contribution of this factor to phenotypic variation in life-history traits. During the reproductive season, we kept female northern grass lizards, Takydromus septentrionalis (Lacertidae), in three temperature-controlled rooms (25, 28 and 32 °C) to measure the effect of the maternal thermal environment on reproductive traits. Maternal thermal environment remarkably affected reproductive frequency and thereby seasonal reproductive output, but had little effect on reproductive traits per clutch or hatchling traits. Females kept at 32 °C produced more clutches and thus had shorter clutch intervals than females from 28 to 25 °C. Clutch size, clutch mass, relative clutch mass, egg size and hatchling traits did not vary among the three treatments. The eggs produced by the females were incubated at 27 °C and the traits of hatchlings were measured. The result that egg (offspring) size was independent of maternal thermal environments is consistent with the prediction of the optimal egg size (offspring) theory. The eggs produced by low temperature females (28 and 25 °C) took longer time to complete their post-oviposition development than did eggs produced by high temperature females (32 °C). This suggests that the eggs from low temperatures might have been laid when the embryos were at relatively early stages. Therefore, maternal thermal environment prior to oviposition could affect post-oviposition development in T. septentrionalis.     

Climate change,boundary increase and elongation of a pre‐existing cline: A case study in Drosophila ananassae

During the past two to three decades, Drosophila ananassae, a warm adapted tropical species, has invaded low to mid altitude localities in the western Himalayas. Due to its cold sensitivity, this species had never been recorded from higher latitudes as well as altitudes in India to the 1960s. A latitudinal cline in this desiccation‐sensitive species corresponds with southern humid tropical localities rather than northern drier subtropical localities. An extension of its cline into lowland to midland montane localities has resulted due to global climatic change as well as local thermal effects through anthropogenic impact. However, D. ananassae populations at species borders are characterized by lower genetic variability for body melanization as well as for desiccation resistance. There is a lack of thermal plastic effects for body melanization, and the observed extended cline might represent evolutionary (genetic) response due to selection pressure imposed by drier habitats. A comparison of fecundity, hatchability and viability at three growth temperatures (17, 20 and 25°C) showed significant reduction in trait values at 17°C in D. ananassae. Thus, its recent range expansion into northern montane localities might involve genetic effects on stress‐related traits and plastic effects on life history traits. We suggest that D. ananassae could serve as an indicator species for analyzing range expansion under changing climatic conditions.     

WITHIN- AND BETWEEN-GENERATION EFFECTS OF TEMPERATURE ON THE MORPHOLOGY AND PHYSIOLOGY OF DROSOPHILA MELANOGASTER

We investigated the effects of developmental and parental temperatures on several physiological and morphological traits of adult Drosophila melanogaster. Flies for the parental generation were raised at either low or moderate temperature (18°C or 25°C) and then mated in the four possible sex-by-parental temperature crosses. Their offspring were raised at either 18°C or 25°C and then scored as adults for morphological (dry body mass, wing size, and abdominal melanization [females only]), physiological (knock-down temperature, and thermal dependence of walking speed), and life history (egg size) traits. The experiment was replicated, and the factorial design allows us to determine whether and how paternal, maternal, and developmental temperatures (as well as offspring sex) influence the various traits. Sex and developmental temperature had major effects on all traits. Females had larger bodies and wings, higher knock-down temperatures, and slower speeds (but similar shaped performance curves) than males. Development at 25°C (versus at 18°C) increased knock-down temperature, increased maximal speed and thermal performance breadth, decreased the optimal temperature for walking, decreased body mass and wing size, reduced abdominal melanization, and reduced egg size. Parental temperatures influenced a few traits, but the effects were generally small relative to those of sex or developmental temperature. Flies whose mother had been raised at 25°C (versus at 18°C) had slightly higher knock-down temperature and smaller body mass. Flies whose father had been raised at 25°C had relatively longer wings. The effects of paternal, maternal, and developmental temperatures sometimes differed in direction. The existence of significant within- and between-generation effects suggests that comparative studies need to standardize thermal environments for at least two generations, that attempts to estimate “field” heritabilities may be unreliable for some traits, and that predictions of short-term evolutionary responses to selection will be difficult.     

Genetic variation and plasticity of thorax length and wing length in Drosophila aldrichi and D. buzzatii

Reaction norms across three temperatures of development were measured for thorax length, wing length and wing length/thorax length ratio for ten isofemale lines from each of two populations of Drosophila aldrichi and D. buzzatii. Means for thorax and wing length in both species were larger at 24 °C than at either 18 °C or 31 °C, with the reduction in size at 18 °C most likely due to a nutritional constraint. Although females were larger than males, the sexes were not different for wing length/thorax length ratio. The plasticity of the traits differed between species and between populations of each species, with genetic variation in plasticity similar for the two species from one locality, but much higher for D. aldrichi from the other. Estimates of heritabilities for D. aldrichi generally were higher at 18 °C and 24 °C than at 31 °C, but for D. buzzatii they were highest at 31 °C, although heritabilities were not significantly different between species at any temperature. Additive genetic variances for D. aldrichi showed trends similar to that for heritability, being highest at 18 °C and decreasing as temperature increased. For D. buzzatii, however, additive genetic variances were lowest at 24 °C. These results are suggestive that genetic variation for body size characters is increased in more stressful environments. Thorax and wing lengths showed significant genetic correlations that were not different between the species, but the genetic correlations between each of these traits and their ratio were significantly different. For D. aldrichi, genetic variation in the wing length/thorax length ratio was due primarily to variation in thorax length, while for D. buzzatii, it was due primarily to variation in wing length. The wing length/thorax length ratio, which is the inverse of wing loading, decreased linearly as temperature increased, and it is suggested that this ratio may be of greater adaptive significance than either of its components.     

SINGLE- AND MULTIPLE-LOCUS GENOTYPES AND LIFE-HISTORY RESPONSES OF GAMBUSIA HOLBROOKI REARED AT TWO TEMPERATURES

Eastern mosquitofish (Gambusia holbrooki) were reared from birth to 10 wk of age at 25°C and 32°C. Relationships of growth, time to maturity, and developmental stability to isozyme genotype were used to examine the hypothesis that more heterozygous individuals should exhibit superior performances, especially under thermally stressful (32°C) conditions. More heterozygous fish grew faster than homozygous individuals, especially at 32°C. Significant differences in time to maturity were detected among allozyme genotypes but not with heterozygosity. Multiple-locus heterozygosity was negatively related to fluctuating asymmetry. Thus, life-history traits were affected by both multiple-locus heterozygosity and single-locus genotype.     

The genetics of phenotypic plasticity. III. Genetic correlations and fluctuating asymmetries

We examined the relationship of three aspects of development, phenotypic plasticity, genetic correlations among traits, and developmental noise, for thorax length, wing length, and number of sternopleural bristles in Drosophila melanogaster. We used 14 lines which had previously been selected on either thorax length or plasticity of thorax length in response to temperature. A half-sib mating design was used and offspring were raised at 19° C or 25° C. We found that genetic correlations were stable across temperatures despite the large levels of plasticity of these traits. Plasticities were correlated among developmentally related traits, thorax and wing length, but not among unrelated traits, lengths and bristle counts. Amount of developmental noise, measured as fluctuating asymmetry and within-environmental variation, was positively correlated with amount of plasticity only for some traits, thorax length and bristle number, and only at one temperature, 25° C.     

Response of Development and Body Mass to Daily Temperature Fluctuations: a Study on <Emphasis Type="Italic">Tribolium castaneum</Emphasis>

Differences in thermal regimes are of paramount importance in insect development. However, experiments that examine trait development under constant temperature conditions may yield less evolutionarily relevant results than those that take naturally occurring temperature fluctuations into account. We investigated the effect of different temperature regimes (constant 30 °C, constant 35 °C, fluctuating with a daily mean of 30 °C, or fluctuating with a daily mean of 35 °C) on sex-specific development time and body mass in Tribolium castaneum. Using a half-sib breeding design, we also examined whether there is any evidence for genotype-by-environment interactions (GEI) for the studied traits. In response to fluctuating temperature regimes, beetles demonstrated reaction norm patterns in which thermal fluctuations influenced traits negatively above the species’ thermal optimum but had little to no effect close to the thermal optimum. Estimated heritabilities of development time were in general low and non-significant. In case of body mass of pupae and adults, despite significant genetic variance, we did not find any GEI due to crossing of reaction norms, both between temperatures and between variability treatments. We have observed a weak tendency towards higher heritabilities of adult and pupa body mass in optimal fluctuating thermal conditions. Thus, we have not found any biasing effect of stable thermal conditions as compared to fluctuating temperatures on the breeding values of heritable body-size traits. Contrary to this we have observed a strong population-wide effect of thermal fluctuations, indicated by the significant temperature-fluctuations interaction in both adult and pupa mass.     

Effect of Temperature on Longevity and Productivity in Drosophila Ananassae: Evidence for Adaptive Plasticity and Trade-Off Between Longevity and Productivity

The genetic response of body size to temperature in the laboratory provides an interesting example of phenotypic plasticity. We found that females of Drosophila ananassae reared to adulthood at 18°C showed significant increase in body weight as compared to females reared at 25°C. At a given temperature, early productivity and lifetime productivity were the highest when the rearing and test temperature were the same. The effect of test temperature was highly significant for total productivity and early productivity. The interaction between test temperature and development temperature was also highly significant. Effect of development temperature was not significant. The females reared at 18°C showed greater body weight but their productivity was not significantly higher than smaller females reared at 25°C. Thus, the usually close relationship between size and fecundity is lost when the size change is due to rearing temperature. These findings provide evidence for adaptive plasticity in D. ananassae. We also found a negative correlation (trade-off) between longevity and productivity, the first report of such a trade-off between longevity and productivity in D. ananassae.     

Effect of temperature on the life history of <Emphasis Type="Italic">Eretmocerus</Emphasis> sp. nr. <Emphasis Type="Italic">furuhashii</Emphasis>, a parasitoid of <Emphasis Type="Italic">Bemisia tabaci</Emphasis>

Eretmocerus sp. nr. furuhashii (Hymenoptera: Aphelinidae) is an indigenous parasitoid of Bemisia tabaci (Gennadius)(Hemiptera: Aleyrodidae) from southern China; the effects of constant temperatures on the life history of E. sp. nr. furuhashii were examined in the laboratory. The developmental period ranged from 39.2 days at 20°C to 12.40 days at 32°C. A total of 263.4 degree-days were required to complete development with a lower developmental threshold temperature of 11.1°C. Of the eggs produced, 59.3% completed development at 20°C with completion increasing to 71.5% at 26°C. Adult female longevity was 10.8 days at 20°C and 5.2 days at 32°C while the mean daily offspring reproduced per female was highest at 29°C with 5.9 offspring. Adult oviposition peaked three days after emergence at 26, 29 and 32°C, and four days post-emergence at 20°C and 23°C. The total numbers of offspring produced per female ranged from 25.7 individuals at 32°C to 41.1 individuals at 20°C. The sex ratio had a female bias and ranged from 0.72 at 17°C to 0.51 at 35°C. The intrinsic rate of increase was 0.1727 at 29°C followed with 0.1606 at 32°C. Results indicated that E. sp. nr. furuhashii reaches its maximum biological potential at temperatures ranging from 26°C to 32°C.     

Age-specific fecundity and life table of Trissolcus semistriatus, an egg parasitoid of the sunn pest Eurygaster integriceps

The effects of temperature on age‐specific fecundity and life table parameters of the egg parasitoid Trissolcus semistriatus (Nees, 1834) (Hymenoptera: Scelionidae) were examined under four constant temperature conditions (17, 20, 26 and 32°C), using eggs of the sunn pest Eurygaster integriceps Puton, 1881 (Hemiptera: Scutelleridae), an important pest of wheat, as hosts. The intrinsic rate of increase increased linearly, while the mean generation time and the doubling time decreased with increases in temperature. The net reproductive rate, however, varied without clear correlation with temperature. Fecundity tended to be higher at higher temperatures. The total number of eggs per female was estimated as 52.0 and 116.4 eggs, respectively, at 17°C and 32°C, with the highest fecundity rate during the first day of oviposition. The oviposition rate fluctuated from 4.4 to 14.3 eggs per day. Oviposition and postoviposition periods and longevity decreased when temperature increased. Maximum longevity for females was 21.6 days at 20°C, and female parasitoids lived longer than males at all temperatures. The development period ranged from 7.1 days (32°C) to 35.6 days (17°C) for males and from 8.4 days (32°C) to 37.2 days (17°C) for females. The development of female T. semistriatus required 166.7 degree‐days (DD) above a theoretical threshold of 11.8°C and the development of males required 142.9 DD above 13.1°C. The numbers of generations per year for female and male T. semistriatus, given the temperature in Tekirdag, Turkey, were estimated to be 9.0 and 8.8, respectively. The potential of the egg parasitoid for the control of E. integriceps is discussed.     

Temperature-dependent development and reproductive traits of <Emphasis Type="Italic">Tetranychus macfarlanei</Emphasis> (Acari: Tetranychidae)

Development and reproductive traits of Tetranychus macfarlanei Baker & Pritchard (Acari: Tetranychidae) were investigated on kidney bean, Phaseolus vulgaris L., at eleven constant temperatures. Tetranychus macfarlanei was able to develop and complete its life cycle at temperatures ranging from 17.5 to 37.5°C. At 15 and 40°C, a few eggs (2–4%) hatched but further development was arrested. Development from egg to adult was slowest at 17.5°C and fastest at 35°C for both females and males. Using Ikemoto and Takai’s linear model, the estimated lower developmental thresholds for egg-to-female adult, egg-to-male adult and egg-to-egg development were 12.9–13.0°C. The thermal constants for the respective stages were 110.85, 115.99 and 125.32 degree-days (DD). The intrinsic optimum temperatures (T _Φ) calculated by non-linear SSI model were determined as 24.4, 24.4 and 24.2°C for egg-to-female adult, egg-to-male adult and egg-to-egg development, respectively. The net reproductive rate (R ₀) was highest at 25°C (167.4 females per female) and lowest at 17.5°C (42.6 females per female). The intrinsic rate of natural increase, r _m, increased linearly with the rising of temperature from 0.102 at 17.5°C to 0.441 day⁻¹ at 35°C. These values suggested that T. macfarlanei could be growing quickly in response to increasing temperatures from 17.5 to 35°C and provide a basis for predicting its potential geographical range.     

Sexual differences in life-history traits in the butterfly Lycaena tityrus: a comparison between direct and diapause development

During direct development the butterfly Lycaena tityrus was previously found to display sex-related reaction norms in response to temperature. Based on selection for protandry in males and fecundity selection for larger females, males favoured early emergence over large size, leading to a dramatic weight loss at higher temperatures, whereas females maintained similar weights throughout. Because males were able to avoid a weight reduction relative to females in spite of their shorter development at lower temperatures, sexual size dimorphism existed at higher temperatures only. In the present paper we compare sexual differences in life-history traits in L. tityrus between direct and diapause development at 25 °C. We demonstrate that, regardless of developmental pathway, protandry persisted and relative sexual size dimorphism, with females being larger, remained unchanged. Although diapausing individuals were less time-constrained, allowing them to grow to considerably higher final weights in both sexes, males were not able to reduce their weight loss relative to females. This is explained by the pressure to gain a developmental advantage solely during post-diapause development, whereas direct developing males may spread the burden over the whole larval period. Our results highlight the importance of considering sexual differences in selective pressures, which may influence central life-history traits in manifold ways.     

ENVIRONMENT-DEPENDENCE OF QUANTITATIVE GENETIC PARAMETERS IN IMPATIENS PALLIDA

Population response to selection depends on the presence of additive genetic variance for traits under selection. When a population enters an alien environment, environment-induced changes in the expression of genetic variance may occur. These could have large effects on the response to selection. To investigate the environment-dependence of genetic variance, we conducted a reciprocal transplant experiment between two ecotypically differentiated populations of Impatiens pallida using the progeny of a standard mating design. The floodplain site was characterized by high water availability, moderate temperatures, and continuous dense stands of Impatiens. The hillside site was drier, with larger temperature extremes and supported only scattered patches of Impatiens with significantly lower seed production and earlier mortality. Estimates of heritability were low for each of the 13 traits measured in each population and site (range from 0–28%). Additive genetic variance for life-history traits tended to be larger than for morphological traits, but genetic variance in fitness was estimated to be not significantly different from zero in all cases. Significant heritability was detected in both populations for one trait (date of first cleistogamous flower) known to be closely related to fitness on the hillside. In general, heritability was reduced for populations when grown in the hillside site relative to the floodplain site, suggesting that stress acts to reduce the expression of genetic variance and the potential to respond to selection there. Consistent reductions in heritability associated with more stressful environments suggest that populations invading such sites may undergo little adaptive differentiation and be more prone to local extinction.     

Developmental temperature responses of Chrysoperla agilis (Neuroptera: Chrysopidae), a member of the European carnea cryptic species group

Chrysoperla agilis Henry et al. is one of the five cryptic species of the carnea group found in Europe. Identification of these species is mainly based on the distinct mating signals produced by both females and males prior to copulation, although there are also morphological traits that can be used to distinguish among different cryptic species. Ecological and physiological cryptic species-specific differences may affect their potential as important biological agents in certain agroecosystems. To understand the effects of temperature on the life-history traits of C. agilis preimaginal development, adult longevity and reproduction were studied at seven temperatures. Temperature affected the development, survival and reproduction of C. agilis. Developmental time ranged from approximately 62 days at 15 °C to 15 days at 30 °C. Survival percentages ranged from 42% at 15 °C to 76% at 27 °C. One linear and five nonlinear models (Briere I, II, Logan 6, Lactin and Taylor) used to model preimaginal development were tested to describe the relationship between temperature and developmental rate. Logan 6 model fitted the data of egg to adult development best according to the criteria adopted for the model evaluation. The predicted lower developmental threshold temperatures were 11.4 °C and 11.8 °C (linear model), whereas the predicted upper threshold temperatures (Logan 6 model) were 36.6 and 36.9 °C for females and males, respectively. Adult life span, preoviposition period and lifetime cumulative oviposition were significantly affected by temperature. The effect of rearing temperature on the demographic parameters is well summarized with the estimated values of the intrinsic rate of increase (r_m) which ranged from 0.0269 at 15 °C to 0.0890 at 32 °C and the highest value recorded at 27 °C (0.1530). These results could be useful in mass rearing C. agilis and predicting its population dynamics in the field.     

Life history responses of the cabbage beetle Colaphellus bowringi to temperature change

Temperature is considered one of the most important mediators of phenotypic plasticity in ectotherms. Here, we investigated life history traits of the cabbage beetle, Colaphellus bowringi Baly (Coleoptera: Chrysomelidae), at a wide range of temperatures (16, 19, 22, 24, 26 and 28°C). The larval and pupal times were significantly decreased with increasing rearing temperature and growth rate was positively correlated with temperature. However, the relationship between body size and rearing temperature in C. bowringi did not follow the temperature–size rule; both males and females reached the highest body weight at 19°C. Females were significantly larger than males at all temperatures. Male pupae lost significantly more weight at metamorphosis compared to females. However, diapausing males gained significantly higher weight after feeding compared to diapausing females at higher temperatures of 22, 24, 26 and 28°C. Body weight tended to decrease with increasing rearing temperature, whereas sexual size dimorphism (SSD) tended to increase with increasing rearing temperature; thus, Rensch's rule is upheld. The degree to which SSD changed with temperature varied with different development stages. SSD was lowest in pupae, highest in newly emerged adults and intermediate in diapausing adults.